Vol. 13(3): 97–103

TERRESTRIAL GASTROPODS OF THE CARPATHIAN BEECH FOREST IN THE MAGURA NATIONAL PARK (SE. POLAND)

KRYSTYNA SZYBIAK1,MA£GORZATA LEŒNIEWSKA1,MA£GORZATA TABORSKA2

1Department of General Zoology, Institute of Environmental Biology, Adam Mickiewicz University, Fredry 10, 61-701 Poznañ, Poland (e-mail: [email protected]), 2Department of Animal Ecology, Institute of Environmental Science, Jagiellonian University, Gronostajowa 7, 30-387 Kraków, Poland

ABSTRACT: The malacocoenosis of rich Carpathian beech forest Dentario glandulosae-Fagetum of the Magura Na- tional Park includes 26 species. The most abundant species in quantitative samples was Carychium tridentatum (Risso), in qualitative samples – Macrogastra latestriata (A. Schmidt). The mean density was 86 indiv. m–2. Domi- nant species were C. tridentatum, Aegopinella pura (Alder) and Vitrea crystallina (O. F. Müller). The species diver- sity index H’ (2.2) and the Pielou index J (30%) were low. The community included mostly typical forest- dwellers. Biogeographically, montane, C. European (lowland and highland), and European components dominated. In its structure the malacocoenosis was much similar to that of the Carpathian beech forest of the Pieniny National Park. Shell parameters in the population of Macrogastra latestriata differed from those found in the Beskid Zachodni and were close to those from the Eastern Carpathian foothills.

KEY WORDS: Magura National Park, Carpathian beech forest, malacocoenosis structure, Macrogastra latestriata

INTRODUCTION

Terrestrial gastropods from the Beskid Zachodni the Magura National Park, has not been studied to Mts have been subject to fairly numerous studies, date. among others by B¥KOWSKI (1878, 1884), KOTULA The aim of the study was an analysis of the struc- (1882), WAGNER (1907), GEYER (1927), URBAÑSKI ture of the malacocoenosis in the Carpathian beech (1932), RIEDEL (1957, 1978), DZIÊCZKOWSKI (1972), forest Dentario glandulosae-Fagetum of the Magura Na- UMIÑSKI (1980) and ALEXANDROWICZ (1984). Most tional Park. Our studies are a part of the project “In- papers, however, contain only faunistic data based ventory and diversity analysis of the Magura National on qualitative samples. The structure of malaco- Park”, and especially of the beech forests of the east- coenoses of the Beskid Niski Mts, and especially of ern end of the Beskid Zachodni Mts.

STUDY AREA

The Magura National Park was established in 1995. rested areas. The park includes two vegetation zones: It is 19,961 ha in area. It is located on the transition the submonate zone up to 530 m a.s.l. occupies about between the Western and Eastern Carpathians, and half of the park area. Fragments of natural forest com- includes the mid part of the Beskid Niski Mts. The munities have been preserved here: oak-hornbeam landscape is formed of low and moderately high hills, forest, Carpathian alder forest, alder swamps, riverine ranging from 380 to 400 m a.s.l.; few exceed 800 m forest. A large area is covered by fir and spruce for- a.s.l. Forests occupy 93% area of the park. The re- ests, as well as plantations with pine as dominant. The maining 7% are meadows, pastures and other non-fo- lower mountain forest zone extends from 530 m a.s.l. 98 Krystyna Szybiak, Ma³gorzata Leœniewska, Ma³gorzata Taborska to the summits. Here rich Carpathian beech forests Preliminary quantitative and qualitative studies on predominate. Small patches of acid beech forest, fir the malacofauna were carried out in the dominant and fir-spruce forest, as well as plantations of birch forest community – the rich Carpathian beech forest and pine, are also found. The vegetation cover of the Dentario glandulosae-Fagetum. It occupies richer habi- park has been well preserved. It is largely natural, tats in lower parts of the slopes. In the herb layer little changed, with a high percentage of protected Dentaria glandulosa and Symphytum cordatum dominate. species (FALIÑSKI 1975, MICHALIK 1981, 1995, DENISIUK et al. 1990).

METHODS

Quantitative studies, carried out in September modified by Horn, Marczewski and Steinhaus similar- 2002, included two localities. Quantitative samples, of ity index, ecological and biogeographical grouping litter and surface layer of soil, were taken with a 25 × (MARCZEWSKI &STEINHAUS 1959, PIELOU 1974, 25 cm frame, to a total of 1 m2 (16 samples) in each ALEKSANDROWICZ 1987, RIEDEL 1988, TROJAN 1992, site. The samples were hand-sorted. Only live speci- GÓRNY &GRÜM 1993, MAGURRAN 1996). The nomen- mens were taken into account in statistical analysis. In clature and systematics follow KERNEY et al. (1983). June and July 2001, gastropods were collected by di- The population of Macrogastra latestriata was subject to rect search. The structure of the malacocoenosis was biometrical analysis. The following shell characters described using the following indices and parameters: were analysed: height, width, aperture height and density, dominance (D), frequency (F), Shannon in- width, number of ribs on the penultimate whorl. The dex of species diversity (H´), species diversity index analysis involved 93 adult, undamaged shells. TDI, Pielou equitability index (J), Morisita index as

RESULTS

The total of 364 specimens, collected in the rich The mean density was 86 indiv. m–2. The most Carpathian beech forest, represented 26 snail species abundantly represented species in quantitative of 10 families (Table 1). The richest in species were samples was Carychium tridentatum (54 indiv. m–2;D– Clausiliidae (9 species), followed by Zonitidae (5) 62.2%, F – 12.3%), and in qualitative samples – and Helicidae (3). Quantitative samples contained Macrogastra latestriata. Aegopinella pura showed the 172 specimens representing 16 species which is 62% highest frequency (F – 21.9%, D – 11.6%) (Table 1). all species.

Table 1. Species composition, dominance (D), frequency (F), number of specimens in the Carpathian beech forest of the Magura NP [ecological groups: 1 – typical forest-dwellers, very rarely found in other habitats, 2 – species living mainly in forests but common also in parks and other shaded habitats, 3 – forest and shade-loving species typical for very humid and sometimes marshy habitats, 7 – euryoecious snails, 8 – species of humid but not marshy habitats of varied degree of shadiness, 9 – snails of very humid, periodically flooded habitats; zoogeographical groups: Ce – Central European low- land and upland, Ep – European, Es – Eurosiberian, Hl – Holarctic, Ma – montane (Alpine and Carpathian), Pl – Palaearctic, Se – S. European]

2 Site1&2(2m) Total number of individuals No. Species D% F% (qualitative and quantitative samples) Ellobiidae 1. Carychium minimum O. F. Müller, 1774 [9, Es] 5.2 3.1 9 2. Carychium tridentatum (Risso, 1826) [8, Ep] 62.2 12.5 107 Cochlicopidae 3. Cochlicopa lubrica (O. F. Müller, 1774) [7, Hl] 0.6 3.1 2 Vertiginidae 4. Columella edentula (Draparnaud, 1805) [8, Hl] 0.6 3.1 1 Enidae 5. Ena obscura (O. F. Müller, 1774) [1, Ep] 0.6 3.1 2 6. Ena montana (Draparnaud, 1801) [1, Ce] 2 Terrestrial gastropods of the Magura National Park 99

2 Site1&2(2m) Total number of individuals No. Species D% F% (qualitative and quantitative samples) Endodontidae 7. Discus rotundatus (O. F. Müller, 1774) [2, Ce] 0.6 3.1 1 Vitrinidae 8. Semilimax semilimax (Férussac, 1802) [1, Ce] 0.6 3.1 1 Zonitidae 9. Vitrea crystallina (O. F. Müller, 1774) [2, Ep] 5.8 12.5 10 10. Vitrea transsylvanica (Clessin, 1877) [1, Ma] 3.5 12.5 6 11. Aegopinella pura (Alder, 1830) [1, Ep] 11.6 21.9 20 12. Nesovitrea hammonis (Ström, 1756) [7, Pl] 3.5 18.8 6 13. Daudebardia brevipes (Draparnaud, 1805) [1, Se] 2.3 6.3 4 Euconulidae 14. Euconulus fulvus (O. F. Müller, 1774) [7, Hl] 0.6 3.1 1 Clausiliidae 15. Cochlodina laminata (Montagu, 1803) [1, Ep] 5 16. Cochlodina orthostoma (Menke, 1830) [1, Ce] 6 17. Macrogastra plicatula (Draparnaud, 1801) [1, Ep] 12 18. Macrogastra latestriata (A. Schmidt, 1857) [1, Ma] 112 19. Macrogastra tumida (Rossmässler, 1836) [3, Ma] 1 20. Clausilia pumila C. Pfeiffer, 1828) [3, Ce] 2 21. Balea biplicata (Montagu, 1803) [2, Ce] 18 22. Balea stabilis (L. Pfeiffer, 1847) [1, Ma] 0.6 3.1 5 23. Vestia gulo (E. A. Bielz, 1859) [3, Ma] 25 Helicidae 24. Perforatella incarnata (O. F. Müller, 1774) [1, Ce] 1.2 6.3 2 25. Perforatella vicina (Rossmässler, 1842) [3, Ma] 3 26. Chilostoma faustinum (Rossmässler, 1835) [1, Ma] 0.6 3.1 1 Total specimens 172 364 Total species 16 26

Fig. 1. Ecological structure of the malacocoenosis from the Carpathian beech forest in the Magura NP – species spectrum; 1–9 – ecological groups, for further explanations see heading to Table 1 100 Krystyna Szybiak, Ma³gorzata Leœniewska, Ma³gorzata Taborska

The dominant species were Carychium tridentatum – superdominant, Aegopinella pura – eudominant, Vitrea crystllina and Carychium minimum – dominants. They constituted 85% of the community (Table 1). The remaining species were subdominants (9%), subrecedents (5%) and recedents (1%). All species found in quantitative samples were accidental, with a frequency up to 25%. In the dominance structure of the malacocoenosis subrecedent species formed the highest proportion. The group included eight spe- cies: Cochlicopa lubrica, Columella edentula, Ena obscura, Discus rotundatus, Semilimax semilimax, Euconulus fulvus, Balea stabilis, Chilostoma faustinum. Their rela- tive abundance did not exceed 1.0%. The species diversity index H’ of the malaco- coenosis was 2.2, Pielou (J) index – 30%. The TDI in- dex, directly comparing the equitability and diversity of the community, irrespective from its component taxa, was 0.593. Gastropods of the malacocoenosis of the beech forest represented six out of nine ecological groups distinguished by ALEXANDROWICZ (1987) (Table 1; Fig. 1). Forest species (group 1) and species living mainly in forests but found also in other habitats (group 3), as well as shade-loving species of very hu- Fig. 2. Zoogeographical composition of the malacocoenosis mid habitats (group 2) formed 76% of the mala- of the Carpathian beech forest in the Magura NP. For cocoenosis (Table 1). The remaining species were further explanations see heading to Table 1

Fig. 3. Shell height in the population of Macrogastra latestriata Terrestrial gastropods of the Magura National Park 101

Fig. 4. Number of ribs on the penultimate whorl in the population of Macrogastra latestriata euryoecious (group 7 – 11%), species of humid habi- ten found on rotting logs and stumps or under tats of varied degree of shadiness (group 8 – 8%) and loose-hanging bark of beech stumps and logs. The species of very humid habitats (group 9 – 5%). shell height ranged from 12.0 to 15.1 mm, the mean The zoogeographical composition of the malaco- was 13.4 mm. The shell width was less variable coenosis is presented in Fig. 2. Montane species (2.9–3.8 mm, mean 3.3 mm). The aperture height formed 26.9%, like C. European lowland and upland and width were 2.4–3.2 mm (mean 2.9) and 2.1–3.0 species (26.9%). The next largest group included Eu- mm (mean 2.4), respectively. The mean number of ropean species (23.1%). Furthermore, Holarctic, ribs on the penultimate whorl was 41 (range 31–55). Palaearctic, S. European and Euro-Siberian element The shell height distribution (Fig. 3) and the distribu- was represented (Table 1). tion of the number of ribs (Fig. 4) in the population The population of Macrogastra latestriata, which was of M. latestriata from the Magura National Park was the most abundant snail in qualitative samples, de- close to normal which suggests a uniformity of the serves a special attention. These snails were most of- population.

DISCUSSION

All the species found in the Carpathian beech for- found under bark of stumps, and on rotting logs, and est of the Magura NP were earlier recorded from the thus absent from the litter samples. The most abun- Beskid Zachodni. Quantitative studies in two sites of dantly represented clausiliid was Macrogastra latestriata. combined area of 2 m2 revealed the occurrence of 16 It constituted 58% of all specimens found during quali- species. Further quantitative studies may well increase tative search. Two members of Clausiliidae: Vestia gulo this number. For comparison, in the Carpathian beech and Balea stabilis, found in the beech forest of the forest of the Pieniny NP (SZYBIAK 2000), quantitative Magura NP., are Carpathian endemics. Cochlodina la- samples from three sites (combined area of 3 m2) minata, C. orthostoma, Macrogastra plicatula, M. latestriata, yielded 42 species, the numbers per site being 28, 21 Balea biplicata and B. stabilis are typical forest-dwellers. and 30. Further ten species were found as a result of di- The remaining three species are clausiliids occurring rect search; eight of them were clausiliids. They were mainly in forests, but typical also of humid habitats: 102 Krystyna Szybiak, Ma³gorzata Leœniewska, Ma³gorzata Taborska

Clausilia pumila, Vestia gulo and Macrogastra tumida.Be- Magura NP is much lower compared to that from the sides the Carpathian endemics, typical Carpathian and Pieniny beech forest (H’ = 4.65, TDI = 0.93). The Boreo-Alpine species are Clausilia pumila and snails of the Magura NP represent the same ecological Macrogastra tumida. Balea stabilis, an E. Carpathian spe- groups as those from the Pieniny, and group 9 – snails cies, occurs in the Magura NP near the western border of very humid habitats. The forest species common to of its distribution range. the two malacocoenoses are: Ena montana, Discus The malacocoenosis of the studied beech forest rotundatus, Vitrea crystallina, V. transsylvanica, has four dominant species. Carychium tridentatum, the Aegopinella pura, Perforatella incarnata, P. vicina, superdominant, and the eudominant Aegopinella pura, Chilostoma faustinum and clausiliids – Cochlodina are common in Europe. C. tridentatum seemed to laminata, C. orthostoma, Macrogastra plicatula, M. show a clustered distribution in the litter, with up to latestriata, M. tumida, Clausilia pumila, Balea biplicata, 37 specimens in some subsamples of 25 × 25 cm. It is a B. stabilis, Vestia gulo. In both areas forest species form very higrophilous species, abundant in litter of wet the same proportion in the malacocoenoses (76% all places. In the Carpathians it occurs at lower altitudes species). Montane species, such as Vitrea transsylva- (BERGER 1963). Its high density and the superdomi- nica, Macrogastra latestriata, M. tumida, Balea stabilis, nant status indicate very favourable conditions for the Vestia gulo, Perforatella vicina and Chilostoma faustinum, species. C. tridentatum is also a dominant (eudomi- found in the Magura NP, were also present in the nant) in the beech forest of the Pieniny NP (SZYBIAK Pieniny beech forest. Their proportion in the two 2000). Aegopinella pura is among dominants in the communities was similar: 26.9% in the Magura NP Polish Carpathian beech forests (RIEDEL 1988); in the and 30.6% in the Pieniny. Pieniny NP it is eudominant (SZYBIAK 2000). The spe- The most abundant clausiliid in the studied cies diversity index H’ of the studied malacocoenosis is malacocoenosis was Macrogastra latestriata. The spe- not high. The low value of the Pielou index (J) is char- cies inhabits mainly beech forests. It is characteristic acteristic of communities in which only few species for the Carpathian lower forest zone. There are nu- dominate. merous records from the Eastern and Western Most species in the malacofauna of the beech forest Carpathians. It was not recorded before from the of the Magura NP are forest-dwellers; the same is true Magurski NP – a transition betwen the Eastern and of other Carpathian beech forests (DZIÊCZKOWSKI Western Carpathians. The shell parameters of speci- 1972, PIECHOCKI &BORCZYK 1990, SZYBIAK 2000). mens from the Magura NP differ from those reported In its species composition the malacocoenosis of by STÊPCZAK (1970) from the Western Beskidy Mts. the Magura NP is much similar to that of the beech The ranges of the shell height and width, as well as ap- forests of the Pieniny NP. The similarity index of erture height in the studied populations are wider, Marczewski-Steinhaus is 0.364. The dominance struc- while the number of ribs on the penultimate whorl is ture of the two malacocoenoses is also rather similar. smaller. In shell size the population is the closest to The Morisita index for the two communities is 0.389. specimens from the foothills of the Eastern Car- The species diversity of the malacocoenosis from the pathians.

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